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\jP Biodiversity 

Proceedings of the Entomological Society of Washington. 

Washington, etc. :Entomological Society of Washington 

v87 1985: 
Page(s): Page 317, Page 318, Page 319, Page 320, Page 321, Page 322 

Contributed by: Smithsonian Institution Libraries 
Sponsored by: Smithsonian 

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87(2), 1985, pp. 317-322 


Beth B. Norden and Suz^anne W. T. Batra 

(BBN) Entomology Department, University of Maryland, College Park, Mary- 
land 20742; (SWTB) Systematic Entomology Introduction Laboratory, Agricul- 
tural Research Service, USDA, Beltsville, Maryland 20705. 

Abstract, — Groups of males of the nearctic Anthophora abrupta Say (Apoidea: 
Anthophorini) chew parsnip tissue, collect the odorous juice in unique adsorptive 
labral mustaches of fine, flattened hairs, and apply it to surrounding objects, 
apparently mixed with their mandibular gland secretion. Such perfumed areas 
outline an oval flight path. This resembles the fragrance-collecting and territorial 
behavior of male neotropical orchid bees (Apidae: Euglossinae). 

The neotropical orchid bees (Apidae: Euglossinae) are well known for the un- 
usual behavior of the males that collect fragrances from flowers and elsewhere. 
Using special pads of adsorptive hairs on their front feet, they brush the surface, 
then pack the collected fragrance into special hair-lined cavities in their hind 
tibiae. These perfumes are evidently used to maintain territories (Dodson, 1973; 
Dressier, 1982). The fragrances may also be sequestered for use in producing male 
mandibular gland attractant pheromones (Williams and Whitten, 1983). Similar 
behavior, not previously known among any other bees, is here reported to occur 
in the nearctic species, Anthophora abrupta Say (Anthophoridae: Anthophorinae). 

A large solitary bee, A. abrupta nests in dense aggregations in vertical clay 
banks or adobe walls (Prison, 1923; Rau, 1929; Norden et al., 1980; Norden, 
1984). Little is known of the sexual behavior of this species. Mating was not seen 
at the nesting site, although occasional males followed and pounced on females 
returning from the field. Caged insects mated on flowers (Norden, 1984). 

For several years, Jean Worthley (pers. comm.) noticed A. abrupta visiting a 
patch of naturalized parsnips at her farm in Owings Mills, Baltimore County, 
Maryland. On the sunny mornings of June 15 and 17, 1982, and June 11, 14, 
and 21, 1984, we visited the site, where we observed and filmed the unique male 
behavior described here. Both years, male A. abrupta were clustering on a parsnip 
plant {Pastinaca sativa L.) growing in partial shade under a loblolly pine about 
75 m from the nest site and 18 m from a small pool where females were ingesting 
water. Neighboring plants that had been previously defoliated by the bees bore 
many brown, necrotic lesions on their stems. Neither male nor female bees were 
visiting the flowers of these parsnips; nor were any bees of either sex seen on 
several cultivated parsnips growing in a sunny garden 50 m away. In 1983, males 
also visited parsnips at the shady site, but not those in the garden (J. Worthley, 
pers. comm.). Males, individually marked while on the parsnip, were observed 
to return to the plant after 1-1.5 hours. 



Fig. L Cluster of four male A. ahrupta chewing on parsnip leaves and petioles. Note portions of 
leaves thai have been shredded bv the bees. 

The male bees alternately chewed the parsnip tissue for 1 1-86 s and periodically 
raised their heads for 2-7 s. Chewing bouts were often terminated when incoming 
bees butted or alighted nearby. Previously chewed areas were preferentially visited. 
When a much-chewed parsnip was removed and replaced with a fresh plant, bees 
visiting the area did not chew on the new plant, but instead located broken, chewed 
sprigs of the old plant lying on the ground. As many as 65 males at a time were 
seen simultaneously chewing on the parsnip's leaves, petioles, or stem (Fig. 1). 
During the chewing phase, the bee's antennae were directed toward the substrate 
(Fig. 2); these normally alert, active bees lost their wariness and could be touched 
by hand. This resembles the oblivious behavior of certain fragrance-collecting 



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; si'fitiiiiv'H 


Figs. 2-5. Behavior of .4. abrupta. 2, Male with deflected antennae, intently chewing on a parsnip 
petiole. 3, Male on bee-damaged leaf, with his head raised and proboscis partly unfolded as he champs 
his mandibles while packing parsnip juice into his labral mustache. 4, Male (arrow) applying parsnip 
odor to grass stem. 5, Female attracted to a netful of females. 

male orchid bees (Dressier, 1982). Males marking objects were much more alert 
than those that were chewing on parsnip, and could not be touched or approached 
within about 30 cm. During the brief head-lifting phase (Fig. 3), the mandibles 
were rapidly vibrated or champed and the proboscis was partly unfolded, appar- 
ently to help drive the plant's juices in among the mustache hairs. 

Males also chewed on the broken ends of dry, dead stems (Fig. 4) in a clump 
of Panicum growing in partial shade about 4 m from the parsnip. Another cluster 
of males was found among grasses in full sun about 30 m from the parsnip. The 
Panicum stems that had been chewed smelled distinctly of parsnip odor; unchewed 
stems had no such odor. When a chewed stem was moved 1 5 cm away, several 
males located and chewed on it. Males also marked two auto bumpers, an Ama- 
ryllis plant, and a honeysuckle bush. They followed the bumpers and Amaryllis 



Figs. 6-10. Labrum and labral hairs o{ Anthophora spp. 6, Labrum of male A. abmpta bearing 
dense mustache ( x 40). 7, Mustachcless labrum of male A. occidentalis as typical of other Anthophorini 
(x40). 8, Densely packed hairs of /i. abmpta mustache (x550). 9, Branched, nonspecialized labral 
hairs of male A. occidentalis ( x 550). 10, Isolated mustache hairs of .4. abrupta ( x 750). These smooth 
hairs have flattened tips that may enhance adsorption by capillarity. 

when they were moved 1-3 m. The group of parsnips, bumpers, Amaryllis, hon- 
eysuckle, and grass locations formed an oval area measuring approximately 1 90 
m X 25 m. This oval area was upwind (prevailing) of most female activity. All 
marked objects were within 1 m of the ground. No females were attracted to the 
males on the parsnip or to those marking elsewhere; however, females (but not 
males) were attracted to other females confined in a net near the parsnips (Fig. 
5), and two females were attracted to a group of netted males taken to the pool. 
Dissection of several males revealed that their mandibular glands produced a 
distinct, sweet fragrance. A similar-smelling, geraniol-like mandibular gland se- 



- ff i *ii S fe> " 






Fig. 1 1. Male A. abrupta marking the rubber bumper of an automobile. The dark patches (arrows) 
are droplets observed to be deposited when males abrade the surface with their mandibles and then 
rub the area with their saturated mustaches. 

cretion of Anthophora occidentalis Cresson attracts other males (Batra, 1978). 
Male A. abrupta are unique in possessing a labral mustache of fine black hairs 
(Figs. 6-10 and Brooks, 1983). This mustache smelled strongly of parsnip odor 
in specimens that were collected as they chewed on the parsnip, bumpers, or grass. 
Some tattered and presumably old males bore mustaches that had a central bald 
area where hairs had evidently worn off during chewing and marking. The spe- 
cialized mustache hairs are densely packed and apically flattened, apparently to 
increase capillary adsorption of the parsnip juice. Certain oil-collecting bees also 
have flattened or otherwise specialized hairs (Vogel, 1971, 1981; Simpson et al., 
1977; Roberts and Vallespir, 1978; Neffand Simpson, 1981; and Dressier, 1982). 
An examination of males and females of 81 species of Anthophorini (76 An- 
thophora, two Hahropoda, two Clisodon, and one Microanthophora) in the col- 
lection of the U.S. National Museum of Natural History revealed no other mus- 
tache-bearing species. Males of 52 species, however, bore sometimes elaborate 
brushes on their mid- and hindtibiae or tarsi that may serve a similar perfume 

gathering and distributing function. Thus, it appears that A, abrupta males are 
unique among Anthophoridae and second only to the Euglossinae in their use of 
plant fragrances in probable combination with mandibular gland pheromones to 
demarcate territories (Fig. 1 1). 

Because parsnip is a recent immigrant of Eurasian origin, the preference of male 
A, abrupta for this plant is puzzling. Perhaps these native bees originally used a 
native species of Umbelliferae. However, they did not visit Daucus or Crypto- 
taenia growing in the garden. Specialized phytophagous insects apparently use 
the toxic furanocoumarins of Umbelliferae as host-recognition cues (Berenbaum, 


1981a). Il is interesting that A. abriipta prefers parsnips growing in shade, which 
produce fewer furanocoumarins than those growing in the sun (Berenbaum, 1981b). 
Rau ( 1 929) observed groups of male A. ahrupta persistently chewing on rusty slag 
at a barren patch of ground about 100 m from a nest site. These males quite 
possibly were marking a territory similar to the one we have observed. 


We thank Jean and Elmer Worthley for calling our attention to the bees visiting 
parsnip, for their observations, and their hospitality. We also thank Norita Cha- 
ncy, Agricultural Research Service, USDA, who provided expert assistance with 
the SEM studies, and Jack Neff, Central Texas Melittological Institute, Austin for 
reviewing the manuscript. 

Literature Cited 

Batra, S. 1978. Aggression, territoriality, mating and nest aggregation of some solitary bees (Hy- 

menoptera: Halictidae, Megachilidae, Collelidae, Anthophoridae). J. Kans. Entomol. See. 51: 

Berenbaum, M. 1981a. Patterns of furanocoumarin production and insect herbivor>' in a population 

of wild parsnip {Pastinaca saliva L.)- Oecologia 49: 236-244. 
. 1981b. Patterns of furanocoumarin distribution and insect herbivory in the Umbelliferae: 

plant cllemistry and communily structure. Ecology 62: 1254-1266. 

Brooks, R. 1983. Systematics and bionomics of A nth ophora: the homboides group and species groups 

of the New Worid. Univ. Calif Publ. Entomol. 98: 1-86. 
Dodson, C. 1973. Co-evolution of orchids and bees. Pp. 91-99. In L. Gilbert and P. Raven, eds,, 

Coevolution of animals and plants. Univ. Texas Press, 246 pp. 
Dressier, R. 1982. Biology of the orchid bees (Euglossini). Ann. Rev. Ecol. Syst. 13: 373-394. 
Prison, T. 1923. Notes on the life history, parasites and inquiline associates of Anthophora abrupta 

Say, with some comparisons with the habits of certain other Anthophorinae (Hymenoptera). 

Trans. Am. Entomol. Soc. 48: 137-156. 
NefT, J. and B. Simpson. 1981. Oil-collecting structures in the Anthophoridae (Hymenoptera): mor- 
phology, function, and use in systematics. J. Kans. Entomol. Soc. 54: 95-123. 
Norden, B. 1984. Nesting biology of Anthophora abrupta (Hymenoptera: Anthophoridae), J. Kans. 

Entomol. Soc. 57: 243-262. 
Norden, B., S. Batra, H. Fales, A. Hefctz, and G. Shaw. 1980. Anthophora bees: unusual glycerides 

from maternal Dufour's glands serve as larval food and cell lining. Science 207: 1095-1097. 
Rau, P. 1929. The biology and behavior of mining bees, Anthophora abrupta and Entechnia taurea. 

Psyche 36: 155-181. 
Roberts, R. and S. Vallespir. 1978. Specialization of hairs bearing pollen and oil on the legs of bees 

(Apoidea: Hymenoptera). Ann. Entomol. Soc. Am. 71: 619-627. 
Simpson, B., J. Neff, and D. Seigler. 1977. Kramena, free fatty acids and oil-collecting bees. Nature 

267: 150-151. 
Vogel, S. 1971. Olproduzierende Blumen, die durch olsammelnde Bienen bestaubt werden. Natur- 

wissenschaften 58: 58-59. 
. 1981. Abdominal oil-mopping — a new type of foraging in bees. Naturwissenschaften 67: 

Williams, N, and W. Whillen. 1983. Orchid floral fragrances and male euglossine bees: methods 

and advances in the last sesquidecade. Biol. Bull. 164: 355-395.